Nomenclatural review of names published in the fungal genus Dermoloma (Basidiomycota, Agaricales, Tricholomataceae) based on morphological analyses of type specimens

Biodiversity Data Journal 13: 158080 CO)
doi: 10.3897/BDJ.13.e158080 open access
Taxonomy & Inventories

Nomenclatural review of names published in the
fungal genus Dermoloma (Basidiomycota,
Agaricales, Tricholomataceae) based on
morphological analyses of type specimens

Michaela Cabonovat, Marisol Sanchez-Garcia§, Miroslav Cabon?, Katarina Adaméikoval, Pierre-
Arthur Moreau!, Alfredo Vizzini*#, Slavomir Adaméik?, Sona Janéovicova"

+ Department of Microbial and Plant Interactions, Plant Science and Biodiversity Centre, Slovak Academy of Sciences,
Bratislava, Slovakia

§ Department of Forest Mycology and Plant Pathology, Swedish University of Agricultural Sciences, Uppsala, Sweden

| Department of Plant Pathology and Mycology, Institute of Forest Ecology, Slovak Academy of Sciences, Zvolen, Slovakia

4 Laboratoire de Génie Civil et géo-Environnement, University of Lille, Lille, France

# Department of Life Sciences and Systems Biology, University of Turin, Turin, Italy

= Department of Botany, Faculty of Natural Sciences, Comenius University Bratislava, Bratislava, Slovakia

Corresponding author: Michaela Cabonova (michaela.cabonova@savba.sk)
Academic editor: Renan Barbosa
Received: 06 May 2025 | Accepted: 25 Jun 2025 | Published: 17 Jul 2025

Citation: Cabonova M, Sanchez-Garcia M, Cabon M, AdamGikova K, Moreau P-A, Vizzini A, Adam¢ik S,
Janéovicova S (2025) Nomenclatural review of names published in the fungal genus Dermoloma
(Basidiomycota, Agaricales, Tricholomataceae) based on morphological analyses of type specimens.
Biodiversity Data Journal 13: €158080. https://doi.org/10.3897/BDJ.13.e158080

Abstract
Background

The majority of members of the fungal genus Dermoloma have been described, based on
morphology and without molecular support. Sequencing most Dermoloma type
specimens has been unsuccessful, probably due to degraded DNA, leaving their
taxonomy primarily reliant on morphological characters. In this study, we re-described
nine Dermoloma types, providing standardised morphological descriptions that include
observations of previously undocumented microscopic structures.

© Cabonhova M etal. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source
are credited.

2 Cabonova M etal

New information

The pileipellis structure of D. hybridum, D. inconspicuum and D. intermedium var.
coniferarum differs strongly from the typical Dermoloma pileipellis and these taxa do not
belong to this genus. Dermoloma atrobrunneum and D. hymenocephalum are distinct
taxa which have not been reported recently. The concept of Dermoloma cuneifolium var.
punctipes is based on the presence of dark spots on the stipe. However, our examination
of the type material reveals that its spores are more consistent with those of D.
atrocinereum, a species that can also exhibit dark dots. Dermoloma longibasidiatum is
likely a synonym of this species as well. The name D. pseudocuneifolium has been
misapplied for a species with amyloid spores, but the type has inamyloid narrow spores
characteristic for D. bellerianum. Dermoloma pragense probably represents a distinct, but
recently unrecorded European species defined by large basidiomata and small spores.
The data presented here are essential for future nomenclatural treatments within
Dermoloma, as current phylogenetic studies suggest the presence of a large number of
undescribed species.

Keywords

pileipellis, morphology, degraded DNA, fungaria, grassland agarics

Introduction

Dermoloma (J.E. Lange) Singer ex Herink (Basidiomycota, Tricholomataceae) is a genus
of agaric fungi defined by small- to medium-sized basidiomata with a collybioid to
tricholomatoid habit, dull grey and brownish colouration on pileus and stipe, lamellae
generally emarginate and adnate in their attachment to stipes and the context with a
distinct farinaceous odour. Microscopically, the genus is characterised by a pluristratous
(multi-layered) hymenidermic pileipellis and the presence of clamp connections on
hyphal septa (Sanchez-Garcia et al. 2021). Dermoloma members are soil fungi growing
in different types of grassland and forest ecosystems. The genus is included in the
indicator group of so-called CHEGD fungi (acronym of taxon names Clavariaceae,
Hygrocybe s.|., Entolomataceae, Geoglossaceae and Dermoloma) which are biotrophic
fungi probably forming an unspecified symbiosis with plants (Cabon et al. 2021).

The previous complex taxonomic study of Dermoloma members, based strictly on
morphology, included eight species in Europe (Contu et al. 2008). Sanchez-Garcia et al.
(2021) published first molecular phylogeny of the genus which included 25 European
and six North American species-rank clades Voto (2022). The recent monographic study
of the genus (AdamCikova et al. 2025) described 30 species from Europe (including 16
new species) and three from North America (all of them new to science) supported by
molecular analysis of six DNA regions. No such comprehensive morphological study
exists for Dermoloma species outside Europe.

Nomenclatural review of names published in the fungal genus Dermoloma (Basidiomycota, ... 3

Amongst 28 valid Dermoloma names at species and lower rank, 22 were published from
Europe and six from the Americas (Sanchez-Garcia et al. 2021). In that study, authors
sequenced 19 Dermoloma types from Europe and North America, of which 10 attempts
were successful. These 10 types correspond to seven Dermoloma species, because two
names are synonyms and one is referring to a fungus from a different genus (AdamCikova
et al. 2025). This study presents descriptions of nine Dermoloma types not described in
Adamcikova et al. (2025) providing standardised and detailed micromorphological data
with comments about their classification and distinguishing characters. These types were
either not successfully sequenced or their sequences did not match recently collected
material from Europe or North America. Older names with insufficient original diagnoses
and sometimes with changing taxonomic concepts are actually hampering progress of
new species descriptions because the process of sorting the taxonomic identity of these
names requires time-consuming loan communication and taxonomic expertise
(Dayarathne et al. 2016). However, overlooking old names when publishing new species
due to relying strictly on sequence data may result in negative consequences (Koukol
and Delgado 2021).

Materials and methods

Sampling and morphological study

The types of nine Dermoloma taxa were loaned and studied (Table 1). Amongst these
types, only D. hymenocephalum and D. intermedium var. coniferarum were successfully
sequenced in a previous study by Sanchez-Garcia et al. (2021).

Table 1.

Studied Dermoloma types with information about the fungaria where they are deposited.

Taxon name Country of the type §Fungarium name Acronym
origin

D. atrobrunneum Trinidad Kew Royal Botanic Gardens, England, UK K (M)

D. cuneifolium var. The Netherlands Naturalis Biodiversity Center, Leiden, The L

punctipes Netherlands

D. hybridum France Conservatoire et Jardin botaniques de la Ville de G

Geneve, Switzerland

D. hymenocephalum USA University of Michigan, Michigan, USA MICH
D. inconspicuum Venezuela Kew Royal Botanic Gardens, England, UK K (M)
D. intermedium var. France Université de Lille, Lille, France LIP
coniferarum

D. longibasidiatum Italy Associazione Micologica Bresadola, Trento, Italy AMB
D. pragense Czechia National Museum, Prague, Czechia PRM

D. pseudocuneifolium France Université de Lille, Lille, France LIP

4 Cabonova M etal

Microscopic structures were examined in a solution of ammoniacal Congo red after a
short treatment in aqueous 10% potassium hydroxide (KOH). The amyloid reaction of
spore walls was assessed in Melzer’s reagent and observed using both the standard
approach with initial observation immediately after mounting the object in the reagent
(Melzer 1924) and also following a prolonged (30 min) incubation (with pre-heating) in
Melzer’s reagent, as described by Vizzini et al. (2020). Pileipellis elements near the
pileus margin and the pileus centre were observed and measured separately.

All microscopic observations followed standards of AdamCcikova et al. (2025) and were
observed under an Olympus CX-41 microscope with an oil-immersion lens at a
magnification of 1000. All drawings of microscopic structures, except for spores, were
made with a camera lucida using an Olympus U-DA drawing attachment at a projection
scale of 2000. Spores on lamella sides not attached to basidia were observed and
illustrated using QuickPHOTO MICRO 3.2 software visualising images captured by a
Promicra 3-3CP camera. Enlarged scanned pictures of spores were used for measuring
with an accuracy of 0.1 um and for preparation of line drawings. Terminology of spore
shapes follows Vellinga (1988). All other elements were measured with an accuracy of
0.5 um. Spores, terminal elements in the pileipellis and caulocystidia were measured 30
times per collection, while basidia and marginal cells at lamellae edges were measured
20 times. The microscopic dimensions in the description are presented as a mean value
plus/minus standard deviation, with minimum or maximum values. Q is the length/width
ratio of Spores.

Molecular analysis of type specimens

We attempted to sequence ITS region or its fragments (ITS1/ITS2) for all analysed
specimens. Molecular workflow (DNA extraction, PCR amplification, Sanger sequencing)
followed AdamCikova et al. (2025). In case of succesful sequencing of ITS region, the

respective sequence was deposited in GenBank (https:/Wwww.ncbi.nim.nih.gov/genbank/)
and accession number is indicated after reference in the Nomenclature section.

Taxon treatments

Dermoloma atrobrunneum (Dennis) Singer ex Bon, 1986

Nomenclature

Dermoloma atrobrunneum (Dennis) Singer ex Bon, Doc. Mycol. 65: 51. 1986.
Basionym: Tricholoma atrobrunneum Dennis, Trans. Brit. Mycol. Soc. 34: 476. 1951.
Material

Holotype:

a. country: Trinidad; municipality: St. Joseph; locationRemarks: solitary on the ground in
bamboo plantation; identifiedBy: R. W. G. Dennis; date!ldentified: 29-09-1949;

Nomenclatural review of names published in the fungal genus Dermoloma (Basidiomycota,... 5

collectionID: Dennis 8 [K(M) 37579]; institutionCode: K (M); occurrencelD:
D07725D6-541C-545B-8535-5601 19D33AAB

Description

Spores (4.7—)4.9—5.0—5.2(—5.3) x (3.6—)3.8—4.0—4.2(—4.4) um; broadly ellipsoid, Q =
(1.16—)1.21-—1.26—1.31(-—1.37); walls amyloid, thin-walled; hilar appendage ca. 0.5
um long. Basidia (16—)18.5—20.6—23(-24) x (4.5—-)5.5-6.2-7 wm; clavate; with 4
sterigmata. Basidioles first cylindrical, then clavate, ca. 4-6 um wide. Marginal cells
(12—)20-—24.7-30(-34) x (2.5—)3—3.4—4(-—5) ym; cylindrical or sometimes fusiform,
apically diverticulate, branched or coralloid. Pileipellis of mainly one, occasionally
two layers of inflated cells; hyphal terminations with brownish parietal pigments,
occasionally with thickened walls up to 1 um, near septa sometimes with incrusted
dark brown pigments. Terminal cells near pileus margin (27—)30—37.8-—45.5(-—55) x
(12.5—)14.5-17.1-19.5(-24.5) um; usually sphaeropedunculate, sometimes
obpyriform; subterminal cells usually narrower and implemented in intricate hyphae
of subpellis. Terminal cells near pileus centre (15—)19-—27.2-35(—46) x (2.5-)5.5-9—
12.5(-18.5) um; usually clavate, sometimes obpyriform, rarely sohaeropedunculate.
Caulocystidia (15—)18.5—28.6-38.5(-—60) x (3—-)3.5—5.2-6.5(-10) um; clavate, rarely
cylindrical, slightly flexuous, apically obtuse, clustered in small ascending fascicules,
sometimes individual and repent; usually thin-walled. Clamp connections present
(Figs 1, 2).

Figure 1. EE

Dermoloma atrobrunneum [K(M) 37579, holotype], details of the type specimen;

a: Original label of the type specimen;

b: Basidiomata of the type collection.

Notes

Dermoloma atrobrunneum was described from Trinidad (Dennis 1951). Sequencing
of the type specimen failed, but its morphology clearly confirmed the presence of

6 Cabonova M etal

amyloid spores and a cellular pileipellis typical for D. subgen. Amylospora Adamc¢ik. It
is the type species of D. sect. Atrobrunnea Contu, which is classified in this subgenus.
The species differs clearly from the other amyloid-spored North American species in
having small, broadly ellipsoid spores, measuring on average 5.0 x 4.0 um, Q = 1.26
and in diverticulate, branched or coralloid, narrow (ca. 3—4 um wide) marginal cells.

OOODIGOOOO0COOO00O

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ncn Tne

(RIAs

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Figure 2. EE

Dermoloma atrobrunneum [K(M) 37579, holotype], microscopic elements. Scale bar = 5 um
for spores and 10 ym for the other elements. a Spores; b Caulocystidia; c Basidia and
basidioles; d Marginal cells; e Pileipellis elements near pileus margin; f Pileipellis elements near
pileus centre.

Dermoloma cuneifolium var. cuneifolium Arnolds, 1992

Nomenclature

Dermoloma cuneifolium var. punctipes Arnolds, Persoonia 14(4): 529. 1992.

Material

Holotype:
a. country: The Netherlands; stateProvince: Limburg; municipality: Wijlre, “Wrakelberg”;
locationRemarks: in poorly developing hayfield on former arable land on steep calcareous
slope with SW exposure; identifiedBy: E. Arnolds; date!dentified: 22.10.1984; collection|D:

Nomenclatural review of names published in the fungal genus Dermoloma (Basidiomycota, ... 7

Arnolds 5337 (L0821553); institutionCode: L; occurrence!D:
722E4591-6CF 3-524B-941B-26C6E8EBC82D

Description

Spores (5.4—)5.6—6.0-6.4(—7.1) x (4.1-)4.2-4.4—4.6(-4.8) um; broadly ellipsoid to
ellipsoid, Q = (1.27—)1.29-1.37—1.44(—1.55); dextrinoid, inamyloid, thick-walled (walls
ca. 0.5 um); hilar appendage ca. 0.6—0.8 um long. Basidia (7.5—)15—-19.6—24(-27) x
(4.5—-)5—6.0-7(—7.5) um; clavate, often thick-walled and dextrinoid; mostly with 4,
occasionally with 2 and rarely with 3 sterigmata. Basidioles first cylindrical, then
clavate, ca. 5-7 um wide. Marginal cells not differentiated, similar to basidioles on
lamellae sides, mixed with dispersed basidia. Pileipellis of mainly one, occasionally
two layers of inflated cells; hyphal terminations with brownish parietal pigments,
occasionally with thickened walls up to 1 um, near septa often with incrusted dark
brown pigments. Terminal cells near pileus margin (17—)25.5—-37.2-48.5(-61) x
(11-)16.5—20.8-—25(—28) wm; sphaeropedunculate, obpyriform or utriform, often
lobate; subterminal cells often lobate, usually narrower and implemented in intricate
hyphae of subpellis. Terminal cells near pileus centre (14.5—)20—28.5-36.5(-43) x
(12.5-)15.5-18.9-22.5(—26.5) wm; usually obpyriform, subglobose or ellipsoid,
occasionally sphaeropedunculate. Caulocystidia (22—)25.5-38.1-50.5(-73) x
(5.5—)7.3—-9.2—11(-13.5) um; clavate or cylindrical, sometimes centrally constricted,
apically obtuse, repent with ascending tips and often clustered; usually thin-walled,
often thick-walled and with incrusted dark brown pigments. Clamp connections
present (Figs 3, 4).

Figure 3. EE

Dermoloma cuneifolium var. punctipes (L0821553, holotype), details of the type specimen. a
Original label of the type specimen; b Basidiomata of the type collection.

Notes

Dermoloma cuneifolium var. punctipes was originally recognised from the type variety
by darker punctuations of the stipe and darker incrusted pigments on the

Cabonova M etal

caulocystidia (Arnolds 1992). We did not observe these characters in any collections
identified as D. cuneifolium from sequence data; thus, D. var. punctipes probably
corresponds to another Dermoloma species. Stipes with darker granulations were
observed in several species with inamyloid spores, for example, D. atrocinereum, but
these darker dots were usually near the stipe base. Sequencing of the type of D. var.
punctipes failed and sequences of recent collections with darker spots on stipes,
identified as D. var. punctipes, resulted in matches with at least three different
species. Spores of the type specimen are on average 6 x 4.4 um large, which is the
best match for D. atrocinereum amongst species with dark punctuations on the stipe.
We did not decide about the taxonomic status of this variety, but it is clear that the
taxonomic concept, based only on the presence of darker dots on the stipe,
corresponds to more than one species.

JdJO0000D0C00G000

MND arsine
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Poe
oon

Figure 4. EE

Dermoloma cuneifolium var. punctipes (L0821553, holotype), microscopic elements. Scale bar
= 5 um for spores and 10 um for the other elements. a Spores; b Caulocystidia; c Basidia and
basidioles; d Pileipellis elements near the pileus margin; e Pileipellis elements near the pileus
centre.

Nomenclatural review of names published in the fungal genus Dermoloma (Basidiomycota, ... 9

Dermoloma hybridum (Kiihner) Bon, 1979

Nomenclature

Dermoloma hybridum (Kuhner) Bon, Bulletin Annuel de la Fédération Centre-Est
d'Histoire Naturelle et de Mycologie 1:14. 1979.

Basionym: Tricholoma hybridum Kthner, Ann. Sci. Franche-Comte 2: 31. 1947.

Material

Holotype:
a. country: France; stateProvince: Doubs; municipality: Besangon; locality: bois d'Avoudrey;
locationRemarks: under fir mixed with deciduous trees; identifiedBy: R. Kuhner;
dateldentified: 16-10-1946; collectionID: GO0126676; institutionCode: G

Description

Spores (5.2—)5.4—5.8-6.2(-6.7) x (3.4—)3.6—-3.9-—4.2(—4.4) um; ellipsoid to narrowly
ellipsoid, Q = (1.36—)1.40—1.49-—1.59(-—1.73); inamyloid, not dextrinoid, thin-walled;
hilar appendage ca. 0.5—1 um long. Basidia and other elements of the hymenium not
well preserved. Pileipellis a cutis of repent hyphae, composed of chains of inflated,
ellipsoid, cylindrical or clavate cells; hyphal terminations thin-walled, sometimes with
brownish granulose pigments. Terminal cells near pileus margin (20—)22—33.4—-44(-
62) x (7-)8—-9.4—11(-13) wm; clavate, cylindrical or ellipsoid, apically obtuse;
subterminal cells usually equal in size and shape. Terminal cells near pileus centre
(20—)25.5-35.0-44.5(-62) x (8.5—)9.5—11.3-13.5(-17) ym; ellipsoid, obpyriform or
cylindrical, rarely clavate. Caulocystidia not observed. Clamp connections not
observed (Figs 5, 6).

Figure 5. EE

Dermoloma hybridum (G00126676, holotype), details of the type specimen. a Original label of
the type specimen; b Basidiomata of the type collection.

Notes

Dermoloma hybridum was described as Tricholoma hybridum by Kuthner (1947) and
defined by a pileus 70-80 mm in diam., context with no odour and a suprapellis

10 Cabonova M etal

(referred in the original description as epicutis) of cylindrical hyphae. Bon (1979)
combined this species in Dermoloma, although these characters clearly contradict the
definition of the genus. Amplification of the barcode ITS region failed, but our study
confirmed that the pileipellis structure is a cutis composed of chains of ellipsoid
inflated cells, which is more typical for other Tricholomataceae members, including
the genus Tricholoma.

ODDIDODDODOOOODID

Figure 6. EE

Dermoloma hybridum (G00126676, holotype), microscopic elements. Scale bar = 5 ym for
spores and 10 yum for the other elements. a Spores; b Pileipellis elements near the pileus
margin; c Pileipellis elements near the pileus centre.

Cc

Dermoloma hymenocephalum Singer, 1962

Nomenclature
Dermoloma hymenocephalum Singer, Sydowia 15(1-6): 142. 1962.
Basionym: Collybia hymenocephala A.H. Sm., Pap. Michigan Acad. Sci. 26: 61. 1941.

Synonyms: = Replaced synonym: Collybia hymenocephala A.H. Sm., Pap. Michigan
Acad. Sci. 26: 61. 1941, nom. lleg.; non Collybia hymenocephala (Speg.) Speg., Syll.
Fung. 5: 242 (1887).

Nomenclatural review of names published in the fungal genus Dermoloma (Basidiomycota, ... 11

= Hydropus hymenocephalus (A.H. Sm.) Redhead, Sydowia 37: 266. 1984.

= Mycena hymenocephala (A.H. Sm.) A.H. Sm., North American Species of Mycena:
385. 1947.

Material

Holotype:
a. country: USA; county: near Dexter; municipality: Silver Lake; identifiedBy: A. H. Smith;
dateldentified: 23-09-1938; collectionID: Smith 11050 (MICH10228); institutionCode:
MICH

Description

Spores (4.8—-)5.4—5.9-6.4(-6.7) x (3.7-)3.9-4.2—4.5(—5.0) um; ellipsoid to narrowly
ellipsoid, Q = (1.24—-)1.31-1.40-1.49(—1.61); amyloid, thin-walled; hilar appendage
ca. 0.5-0.8 um long. Basidia (24.5—)25.2—28.0—30(-34) x (5.5—)6-6.7-7.5(-8) ym;
clavate; with 4 sterigmata. Basidioles first cylindrical, then clavate, ca. 3-6 um wide.
Marginal cells (10—)12.5-15.4—18.5(—21.5) x (3.5—)3.7—4.9-6(—8) tum; mostly clavate
and similar to basidioles on lamellae sides, but shorter, sometimes fusiform and
apically constricted. Pileipellis of mainly one layer of inflated cells; hyphal
terminations with brownish parietal pigments, occasionally with thickened walls up to
0.5 um. Terminal cells near pileus margin (14.5—)20—26.2—32.5(-40) x (7.5—)12-
16.1-20.5(—24) wm; sphaeropedunculate, obpyriform or utriform, rarely clavate;
subterminal cells often lobate, usually narrower and implemented in intricate hyphae
of subpellis. Terminal cells near pileus centre (15—)19—25.2—31.3(-40.5) x (5.5-)9-—
12.4—16(-—21) ym; usually obpyriform, clavate, subglobose or ellipsoid, occasionally
sphaeropedunculate.

Caulocystidia (10.5—)15.5-19.6—-23.5(—27) x (3.5—)4.5—5.8—7(-—8) ym; clavate, rarely
cylindrical, apically obtuse, ascending or erect, in large, dense clusters; thin-walled,
with brownish parietal pigments. Clamp connections present (Figs 7, 8).

Figure 7. EE
Dermoloma hymenocephalum (MICH 10228, holotype), details of the type specimen. a
Original label of the type specimen; b Basidiomata of the type collection.

12

Cabonova M etal

OODIODDOIGIOObd

Cc

sags HM
RON Ciahlde

we

Figure 8. EE

Dermoloma hymenocephalum (MICH 10228, holotype), microscopic elements. Scale bar = 5
um for spores and 10 um for the other elements. a Spores; b Caulocystidia; c Basidia and
basidioles; d Marginal cells; e Pileipellis elements near the pileus margin; f pileipellis elements
near the pileus centre.

Notes

Dermoloma hymenocephalum was originally placed in the genus Collybia by Smith
(1941) and later hesitantly combined in Dermoloma by Singer (1962), a few years
after the introduction of the genus Dermo/oma in Europe (invalidly by Singer (1955),
but later validated by Herink (1958)). However, the original description mentioned
characters typical for the genus Dermoloma, including inflated pedicellate cells in the
pileipellis, amyloid spores, context with farinaceous taste (but indistinct odour), fragile
context etc. Singer (1975) classified the species in D. sect. Atrobrunnea with some
hesitation. Based on his doubts and suggestions, Redhead (1984) combined this
species into the genus Hydropus. Recently, Sanchez-Garcia et al. (2021) confirmed
by the sequencing of the type specimen that this species belongs in Dermoloma.
Dermoloma hymenocephalum is clearly different from the four North American
species present in the phylogeny of Sanchez-Garcia et al. (2021) and it clustered with
four more collections from Smith’s fungarium in the phylogenetic tree. The long
branch in the ITS tree is possibly caused by a low sequence quality of the type
sample and at least three (possibly all five) collections from Smith’s fungarium may

Nomenclatural review of names published in the fungal genus Dermoloma (Basidiomycota, ... 13

represent D. hymenocephalum. It is characterised by shorter spores compared to
other North American Dermoloma species with amyloid spores.

Dermoloma inconspicuum Dennis, 1961

Nomenclature

Dermoloma inconspicuum Dennis, Kew Bull. 15(1): 78. 1961.

Material

Holotype:
a. country: Venezuela; municipality: Caracas; locality: Dpo. Federal, Botanical Garden;
locationRemarks: on bare soil under trees; identifiedBy: R. W. G. Dennis; date!dentified:
03-07-1958; collection|ID: Dennis 1131 [K(M)147991]; institutionCode: K (M)

Description

Spores (4.9-)5.1-5.3-5.6(-5.7) x (2.9-)3.0-3.2-3.4(-3.5) wm; oblong, Q =
(1.54—)1.59-1.67-1.74(—1.84); inamyloid, not dextrinoid, thin-walled, with oil drops or
refringent contents in central part; hilar appendage ca. 0.4 wm long. Basidia
(13-)15.5-17.0-18.5(-—20) x (4.5—)4.8-5.2-5.5(-6) um; clavate or fusiform; with 4
sterigmata. Basidioles first cylindrical, then clavate, ca. 3.5—7 um wide. Marginal cells
not observed, probably not well differentiated. Pileipellis a trichoderm to almost an
hymeniderm, composed of one (or two) layers of mostly clavate cells; hyphal
terminations with dark brown to almost black parietal pigments, sometimes also with
dark incrusted pigments near basal septa, occasionally with thickened walls up to 0.5
um; subpellis of relatively short (ca. 10-30 ym) and narrow (ca. 2.5—5 um) cells.
Terminal cells near pileus margin (16—)19.5—-34.9-50(-—72) x (4-)5.5—7.9-10(-12.5)
um; usually clavate, sometimes fusiform or subcylindrical, apically obtuse or with
moniliform, simple, branched to coralloid appendages; subterminal usually shorter,
sometimes equally wide and cylindrical or slightly ventricose, sometimes narrower
and branched, usually narrower and implemented in intricate hyphae of subpellis.
Terminal cells near pileus centre (16—)19.5—28.3-37(—49) x (6—-)7.5—9.1-10.7(—15.5)
um; mostly clavate, occasionally sphaeropedunculate, ellipsoid or obpyriform,
apically mostly obtuse and rarely with branched appendages. Caulocystidia absent,
stipe surface covered by narrow (ca. 1.5-3 um wide) hyphae with distant septa and
very scarce repent hyphal terminations. Clamp connections inconspicuous or absent.
(Figs 9, 10).

Notes

Dermoloma inconspicuum, described from Venezuela, was the first member of the
genus included in a molecular phylogenetic study and placed close to Lepijota (Pers.)
Gray in the family Agaricaceae Chevall. (Kropp 2008). Previous results of Sanchez-
Garcia et al. (2021) confirmed that the majority of studied Dermoloma species

14

Cabonova M et al

including the type species belong to the family Tricholomataceae and that D.
inconspicuum is not a member of this genus. The species was placed in the genus
Dermoloma, based on a hymenidermic pileipellis composed of relatively narrowly
clavate, 10-12 um wide terminal cells (Dennis 1961). The morphology of the type
specimen also strongly suggests that this is not a member of the genus Dermoloma,
because of relatively narrow, on average only 7-9 ym wide terminal cells in the
pileipellis with frequent appendages which are often branched.

Figure 9. EE

Dermoloma inconspicuum [K(M)147991, holotype], details of the type specimen. a Original
label of the type specimen; b Basidiomata of the type collection.

JOO00000000 0000000

Sse

(a

Figure 10. | doi |

Dermoloma inconspicuum [K(M)147991, holotype], microscopic elements. Scale bar = 5 ym
for spores and 10 um for the other elements. a Spores; b Caulocystidia; c Basidia and
basidioles; d Pileipellis elements near the pileus margin; e Pileipellis elements near the pileus
centre.

Nomenclatural review of names published in the fungal genus Dermoloma (Basidiomycota, ... 15

Dermoloma intermedium var. coniferarum Bon, 1986

. GenBank ITS MW307771 https ://ncbi.nlm.nih.gov/nuccore/MW307771

Nomenclature

Dermoloma intermedium var. coniferarum Bon, Docums Mycol. 65: 51. 1986.

Material

Holotype:
a. country: France; county: Argol; municipality: Lambibi; identifiedBy: J. Mornand;
dateldentified: 28.10.1982; collectionID: Bon 8116 (LIP); institutionCode: LIP;
occurrence!|D: EAF8B396-F 45A-5447-87B8-29E242B9577E

Description

Spores (7—)7.2—7.6—7.9(-8.5) x (4.7—)5.0—5.2—5.5(—5.7) ym; ellipsoid to narrowly
ellipsoid, Q = (1.29-)1.39-1.45-1 .52(-1.58); amyloid, thin-walled; hilar appendage
ca. 0.3-0.8 um long. Basidia (29—)32—35-—38(—40) x (7—)7.5—7.9-8 .5(—9) um; clavate;
with 4 sterigmata. Basidioles first cylindrical or lageniform, then clavate, sometimes
centrally constricted, ca. 3-7 um wide. Marginal cells (16—)22.5-34.4—46(-63) x 3-
3.9—4.5(—5.5) wm; narrowly fusiform, attenuated or subcylindrical, flexuous, often
moniliform, frequently diverticulate or branched (bifurcated or with lateral branches),
often nodulose or lobate, sometimes almost coralloid, thin-walled. Pileipellis an
intricate trichoderm or a transition to a cutis; terminal cells of two types, large,
(22—)27.5—-37.5—47.7(-60) x (10—)11.9-16.8—21.5(-30) wm, inflated, clavate or
obpyriform cells incrusted by thick yellowish-brownish parietal pigments, mixed with
narrower, (24—)28-—36.9—46(—58) x (3—)3.5—4.3—5(-—6) um, mainly coralloid, flexuous,
lobate, moniliform, not incrusted cells. Caulocystidia (27—)31-—40-—48.5(-63) x
(3.5—)4—5.7-7.5(-9) um; clavate or subcylindrical, simple, branched or coralloid,
lobate, moniliform, occasionally inflated, apically obtuse; thin-walled or occasionally
with thickened walls (up to 0.5 um). Clamp connections present (Figs 11, 12).

Notes

Dermoloma intermedium var. coniferarum was proved not to be a member of the
genus Dermoloma. Previous sequencing of type material indicated that it is identical
with Pseudolaccaria pachyphylla (Fr.) Vizzini & Contu (Sanchez-Garcia et al. 2021).
Our morphological observations of the type specimen revealed the presence of
coralloid hyphal terminations in the pileipellis mixed with large, incrusted and inflated
elements, which demonstrated that this taxon has a very different pileipellis structure
compared to Dermoloma. Some authors of this study also misidentified P
pachyphylla in the field as a Dermoloma species, suggesting that the correct
recognition of this species in the field requires some experience. We recognised the
bitter taste and the radial rimulose or squamulose pattern on the pileus surface in the
mature stage or in dry conditions as a useful diagnostic character of P pachyphylla

16

Cabonova M etal

(see also Lavorato et al. (2015)). The pileus surface of Dermoloma species becomes
occasionally cracked when exposed to dry and windy conditions, but the cracking is
irregular and sometimes concentric.

Nom “Pusat, Kae

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DESCRIPTION MACROSCOPIQUE
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Figure 11. EES)

Dermoloma intermedium var. coniferarum [LIP (Bon 8116), holotype], details of the type
specimen, description form.

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i - i a

Figure 12. EE

Dermoloma intermedium var. coniferarum [LIP (Bon 8116), holotype], microscopic elements.
Scale bar = 5 um for spores and 10 um for the other elements. a Spores; b Caulocystidia; c
Basidia and basidioles; d Marginal cells; e Coralloid hyphae in pileipellis near the pileus margin;
f Large inflated incrusted hyphal terminations in pileipellis near the pileus margin.

Nomenclatural review of names published in the fungal genus Dermoloma (Basidiomycota, ... 17

Dermoloma longibasidiatum Contu, Consiglio & Setti, 2008

Nomenclature

Dermoloma longibasidiatum Contu, Consiglio & Setti, Micol. Veg. Medit. 22(2): 110.
2008.

Material

Holotype:
a. country: Italy; stateProvince: Trentino-Alto Adige; county: Pergine; municipality: Susa;
locationRemarks: grassland on margin of mixed forest of Fagus and Larix; identifiedBy:
G. Consiglio, G. Marasca, B. Oss-Emer; dateldentified: 30-10-1993; collection|D:
GC93318 (AMB); institutionCode: AMB

Description

Spores (5.1—)5.3-5.6—-5.9(-6.2) x (3.6—-)4—4.2—4.5(-4.7) um; broadly to narrowly
ellipsoid, Q = (1.21—)1.26—1.33-1.39(-—1.5); inamyloid, not dextrinoid, thin-walled;
hilar appendage ca. 0.5 um long. Basidia (28—)30—32.5-35(—38) x (6.5—)6.7—7.2-
7.7(—8) um; clavate; with 4 sterigmata. Basidioles ellipsoid, cylindrical or clavate, ca.
3.5—6 um wide. Marginal cells not differentiated. Pileipellis composed of two or three
layers of inflated cells; hyphal terminations thin-walled, often with dark incrusted
pigments near basal septa and on_- subterminal cells; subpellis
pseudoparenchymatous, of irregularly shaped, 5-28 um wide elements. Terminal
cells near pileus margin (14—)19-26.8-34.5(-53) x (6—-)11.5-14.6-18(—-22) wm;
obpyriform or sphaeropedunculate, apically obtuse; subterminal usually inflated,
branched or not, obpyriform or subglobose. Terminal cells near pileus centre
(13—)20.5—-29.5—-38.5(—55) x (8—)11.5-19.2—27(-—43) yum; clavate,
sphaeropedunculate, obpyriform or subglobose, often lobate, apically mostly obtuse.
Caulocystidia (20—)27.5-36.4—45.5(-55) x 7.5-9.6—11.5(-16.5) ym; cylindrical to
clavate, apically obtuse; thin-walled, sometimes clustered in tufts. Clamp connections
present (Fig. 13).

Notes

Dermoloma longibasidiatum, described from Italy, was defined morphologically as a
species similar to D. atrocinereum, but distinguished by longer basidia (33-43 wm
long, Contu et al. (2008)). The type sequencing failed, but according to our
morphological examination, the length of the basidia falls within the range of D.
atrocinereum (on average 29-—32.5 um according to our unpublished observations on
collections used in the phylogeny of Sanchez-Garcia et al. (2021)) and we, therefore,
consider D. longibasidiatum to be a later synonym of this species.

18 Cabonova M etal

OOODOODIYDOVVO
: | I

BES
uf

Figure 13. EE

Dermoloma longibasidiatum [AMB (GC93318), holotype], microscopic elements. Scale bar = 5
um for spores and 10 um for the other elements. a Spores; b Caulocystidia; c Basidia and
basidioles; d Pileipellis elements near the pileus margin; e Pileipellis elements near the pileus
centre.

Dermoloma pragense Kubicka, 1975 [described as ‘pragensis’ ]

Nomenclature

Dermoloma pragense Kubiéka, Ceska Mycol. 29: 31. 1975 [described as ‘pragensis’]

Material

Holotype:
a. country: Czechia; municipality: Prague; locality: Kinského sady; locationRemarks: in grass;
identifiedBy: E. Wichansky; dateldentified: 22-06-1965; collectionID: PRM 611173;
institutionCode: PRM

Description

Spores 4.4—4.8-5.1(-5.9) x (3.4—)3.6—-3.8-4.1(—4.7) wm; broadly ellipsoid, Q =
(1.10—)1.17—1.25-1.32(—1.37); amyloid, thin-walled; hilar appendage ca. 0.5-1 ym
long. Basidia ca. 17-20 x 5-6 um; clavate; with 4 sterigmata. Basidioles ellipsoid,

Nomenclatural review of names published in the fungal genus Dermoloma (Basidiomycota, ... 19

cylindrical or clavate, ca. 3-5 ym wide. Marginal cells (16—)22—27.2-32.5(—40) x
(3.5—)5—6.5—-8(—10.5) um, clavate, occasionally subcylindrical or subcapitate, often
lobate, apically usually obtuse, thin walled. Pileipellis composed of two or three
layers of inflated cells; hyphal terminations thin-walled; subpellis
pseudoparenchymatous, of irregularly-shaped, 3-8 um wide elements. Terminal cells
near pileus margin 16—26.8-39.5(—70) x (6—)8.5—13.6—18(—26) ym; obpyriform or
sphaeropedunculate, apically obtuse; subterminal cells usually inflated, branched or
not, obpyriform, ventricose or subcylindrical, sometimes lobate. Terminal cells near
pileus centre 9-18.8-28.5(—42) x (7-)7.5-10.3-13(-—15) um; subglobose, ellipsoid,
obpyriform or sphaeropedunculate, apically obtuse. Caulocystidia (14—-)29-43.1-
57 .5(—70) x (7—)7.5—8.6—10(—10.5) um; clavate, rarely ellipsoid, apically obtuse; thin-
walled or occasionally with thickened walls (up to 1 um). Clamp connections present
(Figs 14, 15).

Figure 14. EES)

Dermoloma pragense (PRM 611173, holotype), details of the type specimen. a Original label
of the type specimen; b Basidioma of the type collection.

Notes

Dermoloma pragense, described from Czechia (former Czechoslovakia), was
Originally recognised by its amyloid and relatively small spores. There was some
nomenclatural confusion about the validity of the name because it was introduced in
the key without a detailed description (Kubicka 1975). However, the diagnostic
characters of the species are described in Latin as “Sporae amyloideae: Sp. 5-6 x
3.5—4.5 pm” and there is a reference to the type specimen (PRM611173), which
complies with the requirements for valid publication (Turland et al. 2018, Art. 39.1).
Bon (1986) later intended to validate the name at varietal rank as D.
pseudocuneifolium var. pragense Bon, but because he selected his own collection as
the type, he published a new name at the variety rank. Ballero and Contu (1988)
combined Bon's variety at species rank and their name is a heterotypic homonym of
Kubicka's name. Our study is based on the type specimen designated by Kubicka
(1975) and previously reported by Svrcek (1966) as D. cuneifolium. The type has very
small spores (on average 4.8 x 3.8 um, Q = 1.25) which agrees only with the spore
dimensions of the Dermoloma collection SAV F-20229 included in phylogeny of
Sanchez-Garcia et al. (2021). However, the holotype collection of D. pragense differs

20 Cabonova M etal

in having much larger basidiomata (pileus 30 mm in diameter, stipe 5-6 mm wide)
according to Svrcek (1966), while SAV F-20229 has pilei 6-8 mm in diameter and
stipes up to 1.5 mm wide. In our opinion, D. pragense may represent a taxon which is
not represented amongst our recent collections.

sie
ain

nang
Figure 15. EE Ce

Dermoloma pragense (PRM 611173, holotype), microscopic elements. Scale bar = 5 um for
spores and 10 um for the other elements. a Spores; b Caulocystidia; c Marginal cells; d
Pileipellis elements near the pileus margin; e Pileipellis elements near the pileus centre.

Dermoloma pseudocuneifolium Herink ex Bon, 1986

Nomenclature

Dermoloma pseudocuneifolium Herink ex Bon, Doc. Mycol. 17(65): 52. 1986.

Earlier invalid name: Dermoloma pseudocuneilfolium Herink, Acta Musei Horti bot.
Bohemiae 1:62. 1958. [nom. inval., without Latin description]

Material

Holotype:
a. country: France; county: Somme; municipality: Saint-Valery-sur-Somme; identifiedBy: M.
Bon; dateldentified: 10-1968; collectionID: Bon 81006 (LIP); institutionCode: LIP

Nomenclatural review of names published in the fungal genus Dermoloma (Basidiomycota, ... 21

Description

Spores (4.2—)4.8—-5.2—5.6(-5.9) x (3-)3.2-3.5-3.9(—4.3) um; ellipsoid to narrowly
ellipsoid, Q = (1.30—)1.37—1.49-—1.62(-—1.83); inamyloid, not dextrinoid, thin-walled,
dispersed also thick-walled and dextrinoid; hilar appendage ca. 0.8-1 wm long.
Basidia (14—-)16.5-18.2—20(-22) x (4.5—-)5-5.7-6.5(-7) wm; clavate; with 2
sterigmata, thin-walled, occasionally also thick-walled and dextrinoid. Basidioles
cylindrical or clavate, ca. 2.5—5.5 um wide. Marginal cells not differentiated. Pileipellis
composed of one or two layers of inflated cells; hyphal terminations thin-walled, often
with dark incrusted pigments near basal septa and on subterminal cells; subpellis
pseudoparenchymatous, of irregularly-shaped, 5-12 um wide elements. Terminal
cells near pileus margin (27—)30—43.8-58(-82) =x (15.5—)18—23.6-—29(—40) um;
obpyriform or sohaeropedunculate, apically obtuse; subterminal ventricose-inflated or
subcylindrical, branched or not. Terminal cells near pileus centre (20—)29-39.3-
49.5(-61) x (12—)13-16.2—19(—23) um; clavate, sohaeropedunculate or obpyriform,
apically obtuse. Caulocystidia (25—)33.5—41.0-—48(-53) x (5.5—)6.5—8.3-10(-11.5)
um; clavate, apically obtuse; thin-walled. Clamp connections absent (Figs 16, 17).

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Figure 16. EES

Dermoloma pseudocuneifolium [LIP (Bon 81006), holotype], details of the type specimen,
original field and micromorphological notes.

Notes

Dermoloma pseudocuneifolium was introduced by Herink (1958) as an invalid name
(no Latin description) and later validated by Bon (1986) who provided a Latin
diagnosis and designated a personal collection as the holotype. Our sequencing of

22 Cabonova M etal

the type was unsuccessful, but the type specimen (a single basidiome) showed
bisporic basidia without clamp connections and inamyloid, narrow spores, on
average 5.2 x 3.5 um, Q = 1.49. These spores are very narrow and clearly match
those of D. bellerianum Bon presented in the phylogeny by Sanchez-Garcia et al.
(2021). However, Bon’s concept was based on a misapplication of D. cuneifolium by
Josserand (1943) and the protologue as well as Bon's notes attached to the type
specimen both describe the spores as amyloid, 7.5—9 x 4—5 um. Such a discrepancy
suggests a confusion on Bon’s part, the origin of which could not be traced; inamyloid
spores of the type specimen are also contrary to the current name used for a species
with amyloid spores (Wilhelm 1992, Arnolds 1993, Arnolds 1995, Contu et al. 2008,
Sanchez-Garcia et al. 2021). Therefore, we here consider ita nomen dubium.

OOODOOQDOQOTOOIDG
J Naga

NGS

mies

ee

Figure 17. EE

Dermoloma pseudocuneifolium [LIP (Bon 81006), holotype], microscopic elements. Scale bar
= 5 um for spores and 10 um for the other elements. a Spores; b Caulocystidia; c Basidia and
basidioles; d Pileipellis elements near the pileus margin; e Pileipellis elements near the pileus
centre.

Discussion

Amongst the 23 validly published Dermoloma names in Europe and North America at
species and lower rank, Sanchez-Garcia et al. (2021) were able to obtain ITS sequences
from type collections of nine taxa prior to this study. Two of these successfully sequenced

Nomenclatural review of names published in the fungal genus Dermoloma (Basidiomycota, ... 23

types are described here and the morphology of the pileipellis structure was congruent
with their classification, strongly supporting the placement of D. hymenocephalum as a
member of the genus Dermoloma and D. intermedium var. coniferarum as a synonym of
Pseudolaccaria pachyphylla. We also provided morphological evidence for excluding D.
hybridum and D. inconspicuum from Dermoloma. The morphology of D. atrobrunneum,
D. hymenocephalum and D. pragense suggested that these names correspond to taxa
without a recent record. The other three studied types, D. cuneifolium var. punctipes, D.
longibasidiatum and D. pseudocuneifolium, are probably synonyms of other previously
published Dermoloma names, but could not be unambiguously assigned to any of them
by morphological observations and original descriptions without DNA data. Molecular
analysis of old hebarium types represent challenges in all aspects of molecular workflow.
It frequently results in highly fragmented DNA coupled with multiple fungal
contaminations and subsequent poor PCR performance with unspecific amplifications
(Forin et al. 2018). In order to increase success in ITS amplification from degraded
samples, development of highly specific PCR primers is often needed (Bradshaw et al.
2022).

There are six other European Dermoloma names whose types were not successfully
sequenced: D. bellerianum, D. fuscobrunneum P.D. Orton, D. intermedium Bon, D.
jJosserandii P.D. Orton, D. magicum Arnolds and D. murinellum E. Horak. Based on
morphological observations of the type specimens, these names were assigned to
phylogenetically defined species (AdamCikova et al. 2025. In order to stabilise these
species concepts, epitypes were selected for each of them amongst recently collected
and sequenced collections.

Amongst 22 validly published European names, two Dermoloma type collections
remained inaccessible to us. A loan request for D. coryleti Singer & Clemencon to F (Field
Museum of Natural History, Chicago, USA) was not successful, but the original diagnosis
(Singer and Clemengon 1971) describes relatively large spores, absence of distinct
odour and field characters which strongly suggest that this species is not a member of the
genus Dermoloma. Dermoloma clavicystis Voto was described recently (Voto 2022), but
since an ITS sequence was made available by the author, the phylogenetic placement of
this species can be identified and it does not require further analyses from our part.
However, this species also needs urgent morphological revision, because its
morphological circumscription is insufficiently brief and is based on the presence of
marginal cells (presented by the author as cheilocystidia) which proved to be present and
well differenciated in the majority of species within D. subgenus Amylospora. The present
study is crucial for an efficient and stable use of the oldest Dermoloma names. Explaining
concepts of older names only documented by brief and incomplete protologues is a good
practice contributing to nomenclatural stability and important for the consolidation of
further scientific finds (Yurkov et al. 2021). This study is important for aiding in the
delimitation of Dermoloma, but sometimes the conclusion about identity of type
specimens has limitations due to low quality of the fungal material and absence of distinct
morphological differences amongst species (AdamCikova et al. 2025).

24 Cabonova M etal

Acknowledgements

The research of the Slovak team was funded by the Slovak Research and Development
Agency project APVV-20-0257 and Scientific Grant Agency of the Ministry of Education,
Science, Research and Sport of the Slovak Republic and the Slovak Academy of
Sciences, grants VEGA 2/0050/22 and VEGA 1/0346/22. Work of M. Cabonova was
supported by SAS Return Project Scheme.

Konstanze Bensch is acknowledged for nomenclatural advice. The following curators of
fungaria are acknowledged for type loans: Gianfranco Medardi (AMB), Philippe Clerc (G),
Bryn Dentinger (K), Pr Regis Courtecuisse and Christophe Lécuru (LIP), Gerard Thijsse
(L), Timothy James (MICH) and Jan Holec (PRM). Gabriela Kozarova is acknowledged
for technical assistance and high-resolution scanning of line drawings.

Author contributions

Study conception and design were prepared by KA, SA and MicC. Micromorphological
observations were performed by SA, KA and SJ. MSG sequenced type collection and
edited sequences. Funding for the study was obtained by SA, SJ, MirC and MicC.
Manuscript draft was prepared by SA, MirC, AV, PAM and MicC. All authors commented
and approved the final manuscript.

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